ENGLISH ABSTRACT: Understanding how divergent selection results in the evolution of reproductive isolation (i.e. speciation) is an important goal in evolutionary biology. Populations of herbivorous insects using different host plant species can experience divergent selection from multiple selective pressures which can rapidly lead to speciation.
Restio leafhoppers are a group of herbivorous insect species occurring within the Cape Floristic Region (CFR) of South Africa. They are specialised on different plant species in the Restionaceae family. Throughout my thesis I investigated how bottom-up (i.e. plant chemistry/morphology of host plant species) and top-down (i.e. predation and competition) factors drive specialisation and divergence in restio leafhoppers. I also investigated interspecific competition as an important determinant of restio leafhopper community structure.
In chapter 2 I quantified host specificity of restio leafhopper species within a local community for 24 months. I found that restio leafhopper species are highly host specific and potentially synchronised with the growth phases of their host plants.
In chapter 3 I used a network metric, modularity, to determine whether host plant partitioning in a restio leafhopper community is non-random (i.e. driven by a deterministic process). This metric allows the identification of the components underlying host plant partitioning (modules). I then performed experiments to show that modules, and therefore host plant partitioning, can mostly be explained by preference and performance relationships (i.e. bottom-up factors).
In chapter 4 I used null models to test whether niche partitioning in restio leafhopper communities is a general pattern across the landscape. I found non-random niche partitioning, which results from strong host specificity, in all investigated restio leafhopper communities. In addition, I performed binary host choice experiments in the presence and absence of interspecific competition, but found no evidence that interspecific competition narrows host preferences. These findings suggest that host specificity, the cause of niche partitioning, is likely shaped over evolutionary time.
Sampling multiple interaction networks across the CFR, in chapter 5, I tested whether restio leafhopper populations are more host specific in species rich communities and regions in the CFR than in species poor communities and regions. I found no positive relationship between restio leafhopper species richness and host specificity at any scale. These findings suggest that specialisation is not driven by interspecific competition.
In chapter 6 I investigated host shifts in Cephalelus uncinatus. C. uncinatus has a broader distribution than any single restio species that it can use; suggesting that host plant related divergence may result from geographic range expansion. I found that allopatric and parapatric populations, but not sympatric individuals, using different host plants have divergent host preferences. I also found evidence for morphological divergence in traits related to predator avoidance in population pairs that exhibit divergent host preferences.
My findings emphasise the importance of both bottom-up and top-down factors, with the exception of interspecific competition, as determinants of specialisation and divergence in restio leafhoppers. I find no evidence that interspecific competition is an important force structuring restio leafhopper communities. Instead, strongly niche partitioned community structure appears to emerge from the speciation process.